683 MycoKeys MycoKeys 120: 231-254 (2025) DOI: 10.3897/mycokeys.120.154233 Research Article Towards a better knowledge and conservation of cryptic macrolichens in Italy: a revision of the genus Cetrelia (Parmeliaceae, Lecanorales, lichenized Ascomycota) Gabriele Gheza’™®, Chiara Vallese’?®, Luca Di Nuzzo'*®, Simona Corneti'’, Renato Benesperi*®, Elisabetta Bianchi*®, Giulia Canali‘, Silvia Del Vecchio'®, Luana Francesconi"™®, Paolo Giordani*®, Pier Luigi Nimis®®, Walter Obermayer®, Chiara Pistocchi', Helmut Mayrhofer®®, Juri Nascimbene’ no ono FP WO DY — BIOME Lab, Department of Biological, Geological and Environmental Sciences, Alma Mater Studiorum, University of Bologna, Bologna, Italy Department of Earth, Environment and Life Sciences (DISTAV), University of Genova, Genova, Italy Department of Biology, University of Firenze, Firenze, Italy Department of Pharmacy, University of Genova, Genova, Italy Department of Life Sciences, University of Trieste, Trieste, Italy Institute of Biology, University of Graz, Graz, Austria Corresponding author: Chiara Vallese (chiara. vallese@unige. it) OPEN @ ACCESS Academic editor: Pradeep Divakar Received: 28 March 2025 Accepted: 29 May 2025 Published: 8 August 2025 Citation: Gheza G, Vallese C, Di Nuzzo L, Corneti S, Benesperi R, Bianchi E, Canali G, Del Vecchio S, Francesconi L, Giordani P, Nimis PL, Obermayer W, Pistocchi C, Mayrhofer H, Nascimbene J (2025) Towards a better knowledge and conservation of cryptic macrolichens in Italy: a revision of the genus Cetrelia (Parmeliaceae, Lecanorales, lichenized Ascomycota). MycoKkeys 120: 231-254. https://doi. org/10.3897/mycokeys.120.154233 Copyright: © Gabriele Gheza et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Abstract Cryptic species are a challenge for conservation since their ambiguous recognition can hinder a reliable evaluation of their distribution and ecology, thus affecting the assessment of their conservation status. Cetrelia W.L. Culb. & C.F. Culb. is a foliose chlorolichen genus with four species in Europe, which represents a good case-study on this issue. All four sorediate Cetrelia species are morphologically very similar and also show a similar ecology. They can be identified by chemical characters related to their distinctive secondary metabolites, whose diagnostic value is also supported by molecular data. In addition, they are overall rare, and therefore virtually endangered, although in previous assessments they were evaluated as “data deficient” due to the scarcity of available data. The few, old literature records in Italy refer almost exclusively to one species (C. olivetorum), which, however, has been shown to be quite rare in other European countries. To better elucidate the actual distribution of the four species in Italy, we carried out a revision of all the available herbarium specimens and checked several new collections from the main centres of distribu- tion. We analysed 320 specimens from 59 sites, confirming the occurrence of all the four species reported from Europe. Cetrelia monachorum is the most widespread, ranging from the Alps to the Apennines and Sardegna. Cetrelia cetrarioides is less widespread, occurring across the Italian Alps. Cetrelia olivetorum is confined to the Eastern Alps and northern Apennines. Cetrelia chicitae is the rarest, being found only in five sites in the Central and Eastern Alps. All the four species dwell in old, moist montane forests dominated by beech and/or conifers and with long ecological conti- nuity, but they show different biogeographical patterns, which should be considered for planning conservation actions. All the sites hosting Cetrelia species, especially those in which more than one species occur, would deserve protection. Key words: Alps, Apennines, Lichens, Mediterranean 231 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Introduction Cryptic species represent a challenge for conservation, since their difficult recognition can hinder a reliable evaluation of their distribution and ecology, which is likely to affect the assessment of their conservation status (Bickford et al. 2007; Trontelj and FiSer 2008; Hending 2025). This is, however, critical to achieve, in order to set the proper management and conservation actions, and also to foster their inclusion in conservation policies. Among less studied organisms, lichens harbor several genera which include cryptic species, and not only among the inconspicuous ones, i.e., those with crustose growth form, but even in more conspicuous ones, such as in macroli- chens, e.g., in the genera Letharia (Altermann et al. 2016), Parmelia (Divakar et al. 2005; Corsie et al. 2019; Crespo et al. 2020), Parmelina (Nunez-Zapata et al. 2011), Usnea (Randlane et al. 2009). The macrolichen genus Cetrelia is a good candidate as a case study in the aforementioned framework, being characterized by morphologically very similar species. Cetrelia was established by Culberson and Culberson (1968) to segre- gate the so-called “parmelioid Cetrariae”, broad-lobed foliose epiphytic species with chlorococcoid photobiont, formerly attributed to the genus Cetraria Ach., but showing distinct morphological, anatomical and chemical features. Unlike the aforementioned macrolichen genera, which were carefully studied in recent de- cades, Cetrelia has been better known for a longer time in general, albeit being locally understudied. To date, the genus includes 18 species distributed world- wide, with the highest diversity reported from Asia (Randlane and Saag 1991; Mark et al. 2019; Farkas et al. 2021). Five morphotypes, based on the regular presence/type of vegetative propagules, apothecia and/or pseudocyphellae, and six chemotypes, based on the regular occurrence of a specific combination of secondary metabolites (alectoronic and a-collatolic acids; microphyllinic acid; olivetoric acid; anziaic acid; perlatolic acid; imbricaric acid), have been recognized within the genus (Culberson and Culberson 1968; Randlane and Saag 1991). In Europe, the genus is represented by four species, i.e., C. cetrarioides (Duby) W.L. Culb. & C.F. Culb., C. chicitae (W.L. Culb.) W.L. Culb. & C.F. Culb., C. monachorum (Zahlbr.) W.L. Culb. & C.F. Culb. and C. olivetorum (Nyl.) W.L. Culb. & C.F. Culb. All belong to the sorediate morphotype, but each one to a dif- ferent chemotype: alectoronic and a-collatolic acids (C. chicitae), olivetoric acid (C. olivetorum), perlatolic acid (C. cetrarioides) and imbricaric acid (C. monacho- rum) (Culberson and Culberson 1968; Randlane and Saag 1991). The European Cetrelia species can be separated by means of fine morphological characters, e.g., position, shape and size of soralia and pseudocyphellae (Obermayer and Mayrhofer 2007; Farkas et al. 2021), which, however, can often be tricky to in- terpret due to high intraspecific variability. Thus, they can be identified with more ease and certainty by using their secondary chemistry, by means of a thin-layer chromatography (Orange et al. 2010), since they produce distinct and well-recognizable compounds as main secondary metabolites (Randlane and Saag 1991; Mark et al. 2019; Farkas et al. 2021). Besides their similar mor- phology, they share also ecological requirements, with different species often growing together in the same sites (Obermayer and Mayrhofer 2007; Howland and Lendemer 2023), which, however, does not explain why some of them are more widespread, whereas others are found more rarely. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 939 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy In the last decade, following the revision by Obermayer and Mayrhofer (2007), a few records of Cetrelia cetrarioides, C. chicitae and C. monachorum were report- ed from some Italian regions (see literature cited by Nimis and Martellos 2025), but all the historical records of Cetrelia lichens in Italy are under the name Cetre- lia olivetorum (or its earlier synonyms; Fig. 1). However, this species has been reported as one of the rarest Cetrelia in other European countries, e.g., Austria (Obermayer and Mayrhofer 2007), Belarus (Bely et al. 2014; Golubkov et al. 2015), Great Britain (Harrold et al. 2009), Hungary (Farkas et al. 2021), Latvia (Degtja- renko and Moisejevs 2020), Lithuania (Kukwa and Motiejinaité 2012), Poland (Kukwa et al. 2012), and the European part of the former Soviet Union (Randlane and Saag 1991). In the light of this updated and more reliable knowledge, it was unlikely that it would be that widespread in Italy, which suggested the need to study the actual distribution of the species belonging to this genus more in depth. The aim of this study is therefore to provide an updated overview of the ge- nus Cetrelia in Italy, resolving the ambiguity left by the older Italian records that was dragged for decades in lichenological literature, and understanding the real distribution of the four species occurring on the national territory. This is crucial also for conservation purposes: the only red list assessment carried out in Italy so far (Nascimbene et al. 2013a) highlighted the poor knowledge of the genus, assigning C. olivetorum to “near threatened” and C. cetrarioides and C. monachorum to “data deficient”. An updated knowledge would allow a more rigorous assessment and provide a stout support for their inclusion in conser- vation policies, not only at the national level, but also in a European perspective. Materials and methods Sampling and identification All the specimens referable to Cetrelia collected in Italy found in the main Europe- an herbaria were retrieved to be revised. In addition, some of the authors carried out new field surveys specifically to contribute to this study with freshly collected material, exploring both localities where Cetrelia species were already reported and new localities where, based on habitat types, the occurrence of these species was likely, or at least possible. To increase the probability of collecting all possible species in the surveyed areas, field sampling was carried out by extensive walks, conducted by experts, covering almost all areas where the species could potential- ly grow. This resulted in the collection of 320 specimens: 46 from 7 public herbaria (Fl, GE, GZU, LD, LI, SI, TSB) and 274 from new field explorations, that are currently housed in 5 private herbaria (Benesperi, Di Nuzzo, Gheza, Nascimbene, Pistocchi). Although morphological differences occur between the four European Cetrelia species, chemical characters are considered as the mostreliable to distinguish them (Obermayer and Mayrhofer 2007; Farkas et al. 2021; Howland and Lendemer 2023); therefore, our specimens were identified based on chemical characters. Thin-layer chromatography (TLC) was carried out with solvents A, B or B’, and C, according to the method by Orange et al. (2010), to study the main secondary metabolites. It is not infrequent for specimens belonging to different species to grow in- termixed; in some cases, mismatches between chemistry and the morphology of what seemed the only thallus revealed the occurrence of lobes of a different species within the same specimen. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 933 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Elevation 3000 2500 2000 1500 1000 500 0kn i | Figure 1. Distribution of the historical records of Cetrelia in Italy, mostly under the name Cetrelia olivetorum (or earlier synonyms). Frl: Friuli Venezia Giulia; Ven: Veneto; TAA: Trentino Alto Adige; Lomb: Lombardia; Piem: Piemonte; VA: Val d’Aostra; Lig: Liguria; Emil: Emilia Romagna; Tosc: Toscana; Marc: Marche; Umb: Umbria; Laz: Lazio; Abr: Abruzzo; Mol: Molise; Camp: Campania; Pugl: Puglia; Bas: Basilicata; Cal: Calabria; Sar: Sardegna; Sic: Sicilia. DNA extraction, PCR amplification and sequencing A subset of the specimens was subject to molecular analysis to compare the resulting phylogenetic tree with results already available in the literature (Mark et al. 2019). DNA corresponding to 61 selected samples was extracted from freshly collected material within 24 months from collection. For each sample, a section was taken from the youngest, most peripheral, viable areas of the thallus lobes. Approximately 5 to 7 mg of this material was mechanically lysed using a Ribolyser. DNA extraction was conducted with the InviSorb® Spin Plant Mini Kit (Invitek Diagnostics) according to the manufacturer's instructions. Three genomic markers were amplified: the entire internal transcribed spac- er region (ITS), partial intergenic spacer region (IGS) from the nuclear ribosom- al cistron, and fragments from a putative single-copy protein-coding genes, MCM7. Undiluted DNA extracts were subjected to PCR (polymerase chain re- actions) using primers ITS1 and ITS4 (White et al. 1990), IGSf and IGSr (Wirtz MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 934 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy et al. 2008), LecMCM/7f-LecMCM7r (Leavitt et al. 2011) in a reaction volume of 50 ul. Amplification of the ITS was carried out with illustra” PuReTag Ready- To-Go™ (Cytiva) while Amplification of IGS and MCM7 was carried out using DreamTag DNA Polymerase (ThermoFisher Scientific). PCR cycling parameters used for amplifying the ITS and IGS regions were: initial denaturation 94 °C for 3 min, followed by 34 cycles of 94°C for 45s, 55 °C for 45 s, 72 °C for 1 min, and final elongation 72 °C for 5 min, after which the reaction was cooled to a constant 4 °C. Following Schmitt et al. (2009) PCR cycling parameters used for amplifying the MCM7 region were: initial denatur- ation 94 °C for 10 min, followed by 38 cycles of 94 °C for 45 s, 56 °C for 50 s, 72 °C for 1 min, and final elongation 72 °C for 5 min. Electrophoresis confirmed the presence of distinct PCR bands, and all suc- cessful products were sent to the Bio-Fab Research laboratory for purification and subsequent Sanger sequencing. Phylogenetic analyses Sequencing reads were manually verified and edited using FinchTV Version 1.5.0 (Geospiza Research team), with low-quality sequences at both ends removed. These sequences were compared with the public sequence database using the BLASTn searches and the megaBLAST algorithm against the NCBI database in order to verify their identity to Cetrelia and check for any contamination. To root the phylogenetic analyses, 3 sequences respectively for ITS, IGS and MCM7 (KX685872; KX685840; KX685799) and corresponding to Xanthoparmelia con- spersa isolate XACO1 were downloaded from GenBank and selected as the out- group, as recommended by Mark et al. (2019) and Divakar et al. (2015). The se- quences were aligned in the program MAFFT v7 (Katoh and Standley 2013) and default parameters (“auto”) for ITS, |GS and MCM7 were selected. Once aligned, the individual marker alignments were concatenated with MEGA11 (Tamura et al. 2021). Bayesian analyses of the concatenated dataset were carried out in Mr- Bayes.v3.3.7a (Huelsenbeck and Ronquist 2001) via the CIPRES Science Gateway (Miller et al. 2010). Best-fit models of nucleotide substitution were determined to be a SYM + G for ITS dataset, TIM2 + G for IGS dataset and K80 + G for MCM/7, by the Akaike Information Criterion using jModelTest 2.1.10 (Kalyaanamoorthy et al. 2017). We performed the Bayesian analysis by integrating our sequences with those of C. monachorum, C. olivetorum, C. chichitae, and C. cetrarioides as utilized by Mark et al. (2019) and available on GenBank (Sayers et al. 2022). We defined a new matrix with 72 taxa and 1668 characters and performed four inde- pendent runs with four chains. Chains were run for 10,000,000 generations each, while parameters and trees were sampled every 1,000" generation. The resulting phylogenetic trees were visualized with FigTree v1.4.4 (Rambaut 2009). Results General results We retrieved and identified or revised a total of 320 specimens from public her- baria (n = 46: 21 TSB, 7 GE, 7 GZU, 6 FI, 2 LI, 2 SI, 1 LD) and new field collections currently housed in private herbaria (n = 274: 183 herb. Nascimbene, 67 herb. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 995 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Gheza, 19 herb. Benesperi, 3 herb. Di Nuzzo, 2 herb. Pistocchi) (Suppl. material 1). These specimens were collected in a total of 58 sites (Suppl. material 2). Despite almost all the old specimens hosted in Italian public herbaria were labelled as “Cetrelia olivetorum”, only 4 of them (2 Fl and 2 TSB) were confirmed as C. olivetorum, the other 21 being referable to C. monachorum (n = 15: 7 TSB, 3 GE, 3 Fl, 2 SI) and C. cetrarioides (n = 6: 3 TSB, 2 GE, 1 Fl). Other specimens were labelled correctly, i.e., C. cetrarioides (n = 8, TSB) and C. monachorum (n = 1, GE), or incorrectly, but not as “C. olivetorum” (i.e., 1 “C. cf. chicitae” GE and 1 “C. cetrarioides” TSB revealed both to be the real C. olivetorum). Specimens hosted in foreign herbaria had been already revised following a TLC-based ap- proach, revealing the occurrence of C. cetrarioides (n = 5: 3 GZU, 1 LD, 1 LI) and C. monachorum (n = 5: 4 GZU, 1 LI). Newly collected specimens, almost all collected and identified expressly for this study, referred to all the four species, with C. monachorum as the most represented one (n = 162: 119 herb. Nascimbene, 21 herb. Gheza, 19 herb. Benesperi, 3 herb. Di Nuzzo), followed by C. cetrarioides (n = 89: 54 herb. Nascimbene, 33 herb. Gheza, 2 herb. Pistocchi); less represented were C. chicitae (n = 16: 10 herb. Gheza, 6 herb. Nascimbene) and C. olivetorum (n = 6: 4 herb. Nascimbene, 2 herb. Gheza). Distribution of Cetrelia species in Italy Cetrelia species were recorded from nine administrative regions (Table 1). Allthe four European species were found in Friuli Venezia Giulia, Veneto, Trentino Alto Adige, and Lombardia. Two species — C. monachorum and C. olivetorums. str. — were found in Liguria and Emilia Romagna, whereas only one species was found in Piemonte (C. cetrarioides), Toscana (C. monachorum), and Sardegna (C. monachorum). Phylogenetic analyses A total of 51 new sequences were amplified for ITS region, 35 for IGS region and 22 for MCM7 region. All the four Cetrelia species analyzed in this study are represent- ed in these sequences (Suppl. material 3). BLAST searches on GenBank revealed some conflicting results between TLC-based and molecular determination in 8 cas- es (Suppl. material 3), due to the aforementioned occurrence of specimens includ- ing mixed lobes of different species. The phylogenetic tree (Suppl. material 4) re- veals a clade structure consistent with the findings of Mark et al. (2019). The nodes corresponding to the four species groups are well-supported. Cetrelia monacho- rum diverges early as a monophyletic group (posterior probability PP = 1), while the second node (PP = 1) positions the C. olivetorum group as a sister to the clade (PP = 1) that confirms the close relationship between C. cetrarioides and C. chicitae. Taxonomy Based on the morphological and chemical comparison of the collected spec- imens, supported by DNA analyses, we provide an updated description of the morphology and chemical profile for each species. Additionally, we summarize information on their distribution, habitat, and phorophytes in Italy, also com- menting their past literature records in the country. An overview of the main diagnostic morphological characteristics is given in Table 2. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 036 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Table 1. Overview of the genus Cetrelia in Italy. Number of specimens (first number) and number of sites (sec- ond number) are reported for each species in each administrative region. Note that the number of sites per region and per species is not necessarily the sum of the number of sites of all the species/regions, since in many sites more than one species occurred. C. cetrarioides | C. chicitae |C. monachorum | C. olivetorum | No. species | No. specimens |_ No. sites Friuli Venezia Giulia 23 1A 4;2 7A Bal & 2:2 4 50 11 Veneto 2a Zaal Hoe ld 3:4 4 105 2 Trentino Alto Adige 27;14 Teal 47 ;8 242. 4 77 16 Lombardia 28 ; 6 O51 1255 a eb 4 a2 8 Piemonte Ba 2 = = = 1 5 2 Liguria = = a, 2 Sel 2 2 Emilia Romagna = 5 D2. Tal 2 3 Toscana = = 2073 = 1 20 3 Sardegna ot a deel = 1 1 1 No. Regions 5 4 8 No. Specimens 108 16 183 12 319 No. Sites 40 5 38 9 58 Table 2. Overview of the main diagnostic morphological characters between the Italian species of Cetrelia, intended for well-developed, typical specimens; intraspecific variability is high, therefore exceptions are often possible. For further details see Obermayer and Mayrhofer (2007). C. cetrarioides C. chicitae C. monachorum C. olivetorum Pseudocyphellae on Small to large, flat or Large, flat; usually also Small, raised, slightly Small, flat or slightly the upper surface of | rarely raised; often lacking | in the central part of the convex; often lacking concave/immersed; often the lobes from the central part of thallus. from the central part of lacking from the central the thallus. the thallus. part of the thallus. Pseudocyphellae on Small to large, flat on Small, raised, slightly Small, raised, slightly Small, raised, slightly the lower surface of | sterile lobes; small, raised, convex, very rare. convex, very rare. convex, very rare. the lobes slightly convex on fertile lobes; generally common. Soredia Fine, 25-35[-40] um. Coarse, [35-]40—55 um. Coarse, [35-40-55 um. | Fine or coarse, 25-55 um. Marginal soralia Smooth, convex, Strongly twisted- Irregular. Smooth, convex, labriform. undulate, with a crenulate labriform. appearance. Overall appearance Fine soredia in smooth, | Coarse soredia in strongly | Coarse soredia in irregular | Fine (but coarser than convex marginal soralia | twisted-undulate marginal marginal soralia and in C. cetrarioides) or and small to large, flat soralia and large, flat raised pseudocyphellae | coarse soredia in smooth, pseudocyphellae onthe | pseudocyphellae generally on the upper surface. convex marginal soralia lower surface of sterile | only on the upper surface. and small, flat to concave lobes. pseudocyphellae on the upper surface. Cetrelia cetrarioides (Duby) W.L. Culb. & C.F. Culb. Description. Thallus foliose, heteromerous, dorsiventral, loosely attached, forming wavy, wide-spreading, usually orbicular, wide patches. Upper surface greenish-grey, lower surface black in the central part to brown at the lobe edg- es. Lobes broad and rounded, up to 20 mm wide, with raised margins. PSeudo- cyphellae on upper surface punctiform, small to rather large, usually not raised, often lacking in the central parts of thallus; pseudocyphellae on lower surface frequently present, at least on some ascending, contorted lobe apices. Soralia MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 237 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy primarily marginal, elongated, usually very smooth and strongly convex, with fine soredia (25-35 um). Lower surface wrinkled, with scattered, simple, black rhizines and a rhizine-free zone along the margin. Apothecia lecanorine, with brown disc. Found fertile only once in Italy (Piemonte: Val Anzasca). Chemistry. Cortex with atranorin (sometimes in low concentrations); me- dulla with perlatolic acid (major), imbricaric acid (minor or absent), anziaic acid (traces or absent). Distribution in Italy. All of the Alps (108 specimens from 40 sites), but more frequent in the central-eastern part: Friuli Venezia Giulia (23 specimens from 11 sites), Veneto (25 specimens from 7 sites), Trentino Alto Adige (27 specimens from 14 sites), Lombardia (28 specimens from 6 sites), Piemonte (5 specimens from 2 sites). Fig. 2. Habitat. Beech, gray alder, coniferous or mixed beech-coniferous forests in the montane belt (400-1450 m a.s.l.). Phorophytes. Abies alba, Acer pseudoplatanus, Corylus avellana, Fagus syl- vatica, Fraxinus excelsior, Larix decidua, Picea abies, Quercus pubescens, Salix appendiculata, Salix caprea, Salix sp., Ulmus sp. Literature. Confirmed citations: Veneto: Obermayer and Mayrhofer (2007), Nascimbene et al. (2010); Trentino Alto Adige: Obermayer and Mayrhofer (2007), Lang et al. (2010), Nascimbene (2014), Nascimbene and Marini (2015), Trindade et al. (2021), Nascimbene et al. (2022); Lombardia: Ghe- za (2019), Ravera et al. (2021). Erroneous citations (specimens belonging to other species): Emilia Romagna: Tretiach et al. (2008) (C. olivetorum), Fariselli et al. (2020) (C. olivetorum). Specimens examined. See Suppl. material 1. Cetrelia chicitae (W.L. Culb.) W.L. Culb. & C.F. Culb. Description. Thallus foliose, heteromerous, dorsiventral, loosely attached, forming wavy, wide-spreading, usually orbicular, wide rosettes. Upper surface glaucous-grey, lower surface black in the central part to brown at the lobe edg- es. Lobes broad and round, up to 20 mm wide, with raised margins. Pseudo- cyphellae punctiform to irregularly-shaped on upper surface, which are rather large and usually not raised, also developed in the older, central parts of thallus; pseudocyphellae on lower surface not developed, or appearing as very small white dots. Soralia primarily marginal on strongly twisted lobes, giving the lobe- ends a somewhat nibbled appearance, usually smooth, with coarse soredia (35-55 um). Lower surface rather regularly ridged, with scattered, simple, black rhizines and a rhizine-free zone along the margin. Never found fertile in Italy. Chemistry. Cortex with atranorin (sometimes in low concentrations); medul- la with alectoronic, a-collatolic and physodic acids. Distribution in Italy. Central-eastern Alps (16 specimens from 5 sites): Friuli Vene- zia Giulia (4 specimens from 2 sites), Veneto (2 specimens from 1 sites), Trentino Alto Adige (1 specimen from 1 site), Lombardia (9 specimens from 1 site). Fig. 2. Habitat. Beech, coniferous or mixed beech-coniferous forests in the mon- tane belt (870-1195 maz.s.l.), moist due to frequent rainfall and/or fog, usually near streams. Phorophytes. Abies alba, Fagus sylvatica. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 038 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Cetrelia cetrarioides Cetrelia chicitae Elevation 3000 2500 ie 200kin =i 2000 Cetrelia monachorum Cetrelia olivetorum 1500 1000 - Figure 2. Updated distribution of the four Cetrelia species in Italy. Literature. Confirmed citations: Friuli Venezia Giulia: Nascimbene et al. (2021); Veneto: Nascimbene et al. (2021). Remarks. This is the rarest Cetrelia species in Italy, which is in accord with the data from the rest of Europe (Obermayer and Mayrhofer 2007; Harrold et al. 2009). It is instead one of the most widespread species in other areas, e.g., North America (Howland and Lendemer 2023) and India (Mishra and Upreti 2015). Specimens examined. See Suppl. material 1. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 939 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Cetrelia monachorum (Zahlbr.) W.L. Culb. & C.F. Culb. Description. Thallus foliose, heteromerous, dorsiventral, loosely attached, form- ing wavy, wide-spreading, usually orbicular, wide rosettes. Upper surface green- ish-grey, lower surface black in the central part to brown at the lobe edges. Lobes broad and round, up to 20 mm wide, with raised margins. Pseudocyphellae on upper surface frequently developing on slightly raised structures, often appear- ing as accumulation of individuals to form a larger unit, sometimes lacking in the central parts of older thalli; absent from lower surface. Soralia primarily marginal, often rather irregularly shaped, with coarse soredia (35-55 um). Lower surface wrinkled, with scattered, simple, black rhizines and a rhizine-free zone along the margin. Apothecia lecanorine, with brown disc. Found fertile three times in Italy (Trentino Alto Adige: Val Brenta, 1; Veneto: Cansiglio, 2). Chemistry. Cortex with atranorin (sometimes in low concentrations); medul- la with imbricaric acid (major), perlatolic acid (minor or absent), anziaic acid (traces or absent). Distribution in Italy. Central-eastern Alps (155 specimens from 30 sites): Fri- uli Venezia Giulia (21 specimens from 6 sites), Veneto (75 specimens from 11 sites), Trentino Alto Adige (47 specimens from 8 sites), Lombardia (12 speci- mens from 5 sites); Northern Apennines (27 specimens from 7 sites): Liguria (5 specimens from 2 sites), Emilia Romagna (2 specimens from 2 sites), Toscana (20 specimens from 3 sites); Sardegna (1 specimen from 1 site). Fig. 2. Habitat. Beech, coniferous or mixed beech-coniferous forests and chestnut groves from the hilly to the montane belt (274-1600 m a.s.I.). Phorophytes. Abies alba, Acer pseudoplatanus, Alnus incana, Castanea sa- tiva, Corylus avellana, Fagus sylvatica, Fraxinus excelsior, Picea abies, Prunus avium, Quercus cerris, Salix caprea, Salix sp. Literature. Confirmed citations: Friuli Venezia Giulia: Obermayer and Mayrhofer (2007); Veneto: Nascimbene et al. (2021); Trentino Alto Adige: Ober- mayer and Mayrhofer (2007), Nascimbene (2014), Nascimbene and Marini (2015), Trindade et al. (2021), Nascimbene et al. (2022); Liguria: Ravera et al. (2019); Sardegna: Nascimbene et al. (2021). Remarks. The record of “C. olivetorum” by Brackel (2015) should be referred to this species, since the only Cetrelia species found by us in the same site was C. monachorum. Specimens examined. See Suppl. material 1. Cetrelia olivetorum (Nyl.) W.L. Culb. & C.F. Culb. Description. Thallus foliose, heteromerous, dorsiventral, loosely attached, forming wavy, wide-spreading, usually orbicular, wide rosettes. Upper surface greenish-gray to glaucous-gray, lower surface black in the central part to brown at the lobe edges. Lobes broad and round, to 20 mm wide, with raised mar- gins. Pseudocyphellae punctiform on upper surface, which are rather large and usually not raised, also developed in the older, central parts of thallus; pseudo- cyphellae on the lower surface not developed, or appearing as very small white dots. Soralia primarily marginal on strongly twisted lobes, giving the lobe-ends MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 940 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy a somewhat nibbled appearance, usually smooth, with coarse soredia (25-55 um). Lower surface rather regularly ridged, with scattered, simple, black rhizines and a rhizine-free zone along the margin. Never found fertile in Italy. Chemistry. Cortex with atranorin (sometimes in low concentrations); medul- la with olivetoric acid. Distribution in Italy. Central-eastern Alps (10 specimens from 7 sites): Friuli Venezia Giulia (2 specimens from 2 sites), Veneto (3 specimens from 1 site), Trentino Alto Adige (2 specimens from 2 sites), Lombardia (3 specimens from 2 sites); Northern Apennines (2 specimens from 2 sites): Liguria (1 specimen from 1 site), Emilia Romagna (1 specimen from 1 site). Fig. 2. Habitat. Beech or mixed beech-coniferous forests in the montane belt (650- 1450 maz.s.I.), moist due to frequent rainfall and/or fog, often near waterbodies (streams, lakes). Phorophytes. Abies alba, Acer pseudoplatanus, Castanea sativa, Fagus syl- vatica, Picea abies. Literature. Confirmed citations: Friuli Venezia Giulia: Nimis (1982). Dubi- ous citations (specimens not found): Friuli Venezia Giulia: Glowacki (1874), Clerc (1984), Carvalho (1997); Veneto: Du Rietz (1924), Lazzarin (1997), Cani- glia et al. (1999), Thor and Nascimbene (2007), Brackel (2013); Trentino Alto Adige: Lettau (1957), Caniglia et al. (1988), Nascimbene and Caniglia (2000), Nascimbene (2005a, 2006, 2008a), Stofer (2006), Nascimbene et al. (2007, 2022), Lang et al. (2010), Nimis et al. (2015); Lombardia: Stizenberger (1882), Du Rietz (1924); Piemonte: Jatta (1909/11), Martel (1910), Montacchini and Piervittori (1979), Griselli et al. (2003); Emilia Romagna: Zanfrognini (1902), Nimis (1984, 1985), Fariselli et al. (2020); Toscana: Baglietto (1871), Sam- bo (1927); Abruzzo: Jatta (1909/11), Recchia and Villa (1996), Stofer (2006), Gheza et al. (2021); Puglia: Nimis and Tretiach (1999); Sicilia: Marino and Paratore (1900), Jatta (1909/11), Grillo and Caniglia (2004, 2006). Erroneous citations (specimens belonging to other species): Friuli Venezia Giulia: Nimi and Loi (1984) (C. monachorum), Tretiach and Molaro (2007) (C. cetrarioides); Veneto: Nascimbene and Caniglia (1997, 2002, 2003), Nascimbene (2005b, 2008b, 2011), Nascimbene et al. (2005, 2006, 2007, 2009, 2010, 2013b) (all C. cetrarioides and C. monachorum); Trentino Alto Adige: Kernstock (1890) (C. monachorum); Lombardia: Jatta (1909/11) (C. monachorum); Piemonte: Baglietto and Carestia (1865, 1880) (C. cetrarioides), Isocrono et al. (2004) (C. cetrarioides); Liguria: Putorti et al. (1999) (C. monachorum), Giordani and Brunialti (2000) (C. monachorum), Brunialti et al. (2001) (C. monachorum); Toscana: Brackel (2015) (C. monachorum). Remarks. We were not able to retrieve duplicates of the exsiccata distrib- uted by Anzi as Lichenes Rariores Langobardiae n. 48 and Lichenes Minus Rari Italiae Superioris n. 99, cited by Stizenberger (1882) and Du Rietz (1924). The record by Kernstock (1890) likely refers to specimens distributed as Flora Exsiccata Austro-Hungarica n. 3117, of which we revised 5 specimens (GB- 0178237, LD-1086559, and three unnumbered specimens from GZU), all refer- ring to C. monachorum. About the records of “C. olivetorum” by Baglietto and Carestia (1865, 1880), that were cited also by Isocrono et al. (2004), we were able to retrieve just one exsiccatum (LD-1064112) that refers to C. cetrarioides. Specimens examined. See Suppl. material 1. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 241 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Discussion All the four European species of Cetrelia currently occur in Italy, but with differ- ent ranges. The most widespread species is C. monachorum, which was found in 38 sites, ranging from the Alps to the Apennines, and occurring also in one site in Sardegna. C. cetrarioides is widespread as well, occurring in 40 sites, but being limited to the Alps. C. olivetorum shows scattered occurrences in the central-eastern Alps and the northern Apennines, in a total of nine sites. Finally, C. chicitae is confined to a few very well-preserved moist coniferous-beech for- est stands in five sites in the central-eastern Italian Alps. Only three sites, i.e., Pista degli Abeti in Val di Scalve (Lombardia), Foresta del Cansiglio (Veneto) and Lago di Sauris (Friuli Venezia Giulia), hosted all the four Cetrelia species together, whereas five sites (Foresta di Tarvisio and Val Fleons in Friuli Venezia Giulia; Lago di Tovel, Val Brenta and Val Canali in Tren- tino Alto Adige) hosted three species. In all the other 50 sites, only either two or one species were found. The three sites hosting all the four Cetrelia species are remarkable for their high environmental quality and ecological continuity, which reflect in the high number of other rare and/or threatened macrolichens occurring there (Suppl. material 5). This is in accord with what has been ob- served also in North America (Howland and Lendemer 2023). Cetrelia chici- tae, in particular, has always been found related to high-quality forest habitats (Obermayer and Mayrhofer 2007; Howland and Lendemer 2023). In a very few sites we collected fertile material of C. cetrarioides (one specimen in one site in Piemonte) and C. monachorum (one specimen in one site in Trentino Alto Adige and two specimens in one site in Veneto). Fertile specimens have been reported very rarely in contemporary literature (Obermayer and Mayrhofer 2007), and apothecia are likely produced only in highly optimal habitat conditions. Most of our records of all the four Italian Cetrelia species are from well-pre- served, moist forests dominated by conifers and/or beech, located in the montane belt of the Alps and Apennines. Their ecology seems similar, as observed also in other geographical areas (Obermayer and Mayrhofer 2007; Kukwa et al. 2012; Farkas et al. 2021), but differences in distribution suggest that there are also slight differences in ecological preferences, e.g., some species perhaps need more moisture. Cetrelia chicitae, the rarest species, occurs only in very well-preserved forests, in some cases even managed (i.e., Pista degli Abeti and Cansiglio), but granting very long ecological con- tinuity and habitat stability, since the persistence of such forests has been documented for at least three centuries. The occurrence of Cetrelia species, namely C. monachorum and C. olivetorum, in the Northern Apennines seems to be restricted to chestnut groves in the lower montane belt. In that area, this anthropogenic forest type occurs in moist sites and is characterized by a long ecological continuity, also hosting interesting epiphytic assemblages rich in species of conservation concern (Matteucci et al. 2012). The confusion in Italian herbarium material previous to this revision, with most of the specimens misidentified or placed under the rough name of “Cetre- lia olivetorum”, also after the recognition that different chemotypes belong to different species (Obermayer and Mayrhofer 2007), suggests that morphology MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 942 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy and spot test results alone are often not diagnostic enough to identify Cetrelia specimens at the species level with a satisfying degree of certainty. Therefore, TLC should always be carried out when dealing with this genus, since second- ary chemistry is always diagnostic and backed by molecular data (Obermayer and Mayrhofer 2007; Mark et al. 2019; Farkas et al. 2021; Howland and Len- demer 2023). A few morphological characters can be diagnostic in well-de- veloped specimens (Table 2; see also the key by Obermayer and Mayrhofer 2007). Well-developed pseudocyphellae on upwardly curved marginal ar- eas of the underside of the thallus, in combination with smooth soralia, can be found only in C. cetrarioides. Strongly twisted soralia, in combination with very coarse soredia and big, flat pseudocyphellae, only in C. chicitae. Distinctly raised pseudocyphellae, in combination with very coarse-grained soralia, only in C. monachorum. Finally, a C+ clearly red reaction of the medulla, in combina- tion with smooth soralia, is found only in C. olivetorum. Further elements that contribute to complicate the assessment of the actu- al rarity of Cetrelia species are the slightly different microhabitat preferences, which were observed in the field on some occasions. For example, C. chicitae was found very rarely on trunks, more often on branches and twigs; the real distribution of species limited to, or preferentially growing in the canopy can be very difficult to assess reliably (Rosso et al. 2000). Red list assessment in particular risks to be negatively affected by the missed recognition of the species. The preliminary Red List of Italian epi- phytic lichens (Nascimbene et al. 2013a), carried out with a simplified ap- proach, included the three species reported from Italy at the time. Unfortu- nately, the assessment of C. olivetorum (evaluated as “Near-threatened”) was affected by the amount of erroneous literature records, whereas the assessment of C. cetrarioides and C. monachorum, reported from Italy since a few years, was not possible due to the scarcity of data, and both were eval- uated as “Data Deficient”. A red list assessment has not been undertaken in the present work, mainly due to the impossibility of making a comparison with reliable historical data. Other European countries were able to assess Cetrelia species according to the IUCN criteria. In Estonia, C. cetrarioides and C. olivetorum were evaluated as “Endangered”, and C. monachorum as “Critically Endangered” (L6hmus et al. 2019). In Latvia, C. cetrarioides and C. olivetorum were evaluated as “Vulnerable”, and C. monachorum as “En- dangered” (Degtjarenko et al. 2024). In general, the trend seems to indicate that these species are indeed at risk in Europe. Unfortunately, we were not able to verify several historical literature records from southern regions, i.e., those from Abruzzo (Jatta 1909-1911; Recchia and Villa 1996; Stofer 2006), Puglia (Nimis and Tretiach 1999) and Sicilia (Marino and Paratore 1900; Jatta 1909-1911; Grillo and Caniglia 2004, 2006), since specimens referred to those records were not retrieved from any herbarium. Albeit at least some of those records seem reliable, provided the caveat of considering them as “Cetrelia sp.” and not “Cetrelia olivetorum s. str.”, they are mostly old, therefore making the current occurrence of Cetrelia spp. in southern Italy at least dubious. The fact that Cetrelia specimens were not found even in potentially suitable biotopes visited recently by some of us, e.g., the Abetina di Rosello (Nascimbene and Pellegrini 2021) in Abruzzo, support this theory. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 943 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Conclusions All the four Cetrelia species reported from Italy and Europe so far occur in old, moist forests dominated by beech and/or conifers in the montane belt, but they show different biogeographical patterns. Two of them, i.e., C. cetrarioides and C. monachorum, are distributed more widely and are more abundant in their occurrence sites, whereas the other two, i.e., C. chi- citae and C. olivetorum, have a less broad and more scattered distribution and are usually not as abundant as the previous ones. Furthermore, two species, i.e., C. cetrarioides and C. chicitae, are limited to the Alps, whereas the other two, i.e., C. monachorum and C. olivetorum, reach also the northern Apennines. Protection should be advisable for the sites hosting all the four species together, also in consideration of the other species of conservation interest co-occurring with them in those sites. All this should be carefully considered when planning both forest management and conservation ac- tions aimed at including lichens at the local and the national scales. Elucidating the real current distribution of Cetrelia species in Italy is a first step to better understand their biogeography and their ecology, which is needed to include them in conservation policies. In this context, we em- phasize the crucial role of both herbarium collections and targeted field- work in assessing species distribution. Herbarium collections not only can provide a starting point for selecting survey sites, but also allow for direct comparisons between newly collected samples and historical specimens. This approach provides a more comprehensive and reliable distribution assessment than one based solely on field sampling. On the other hand, new fieldwork is crucial, not only to assess the permanence of species in historically known sites, but also to discover new ones and to collect fresh specimens suitable for molecular analyses, which are often not possible on old herbarium material. Accurate knowledge of species distribution forms the foundation for more ap- plied tasks, and represents a fundamental step, especially for cryptic species, for which obtaining reliable and comprehensive data is challenging compared to more easily identifiable ones. Without a solid grasp of their distribution, con- servation strategies may miss vital elements of ecosystem health and biodiver- sity protection. Also, the protection of cryptic species is challenging, due to the difficulty - often the impossibility — to recognize them at the species level with certainty in the field. While waiting for new techniques to be developed to achieve this, a holistic approach to wide-scale protection and conservation, e.g., based on protected areas networks and area-based conservation, is critical to guarantee the survival of such peculiar components of biodiversity (Hending 2025). Acknowledgements We are grateful to: Elena Betti, Eleonora Cominato, Maurizio Salvadori, for providing some specimens; Dr. Ilaria Bonini, Dr. Chiara Nepi, Prof. Mauro Tretiach, for allowing the consultation of specimens from SI, Fl and TSB, re- spectively; Prof. Manuela Mandrone, Prof. Ferruccio Poli, Prof. Lucia Conte, for helping with TLCs. MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 oA Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Funding No funding was reported. Author contributions Conceptualization: GG, CV, LDN, RB, PG, JN. Data curation: GG, CV, LDN, LF, CP. Formal analysis: CV, LDN. Funding acquisition: JN. Investigation: GG, CV, LDN. Methodology: CV, LDN, SC. Project administration: GG. Resources: GG, LDN, RB, PG, CP, JN. Supervision: PLN, WO, HM, JN. Visualization: CV, LDN. Writing - original draft: GG, CV, LDN, SC. Writing - review and editing: RB, EB, GC, SDV, LF, PG, PLN, WO, CP, HM, JN. Author ORCIDs Gabriele Gheza © https://orcid.org/0009-0009-7524-8522 Chiara Vallese © https://orcid.org/0000-0002-8531-5954 Luca Di Nuzzo ® https://orcid.org/0000-0002-5558-9401 Renato Benesperi © https://orcid.org/0000-0003-4296-3393 Elisabetta Bianchi © https://orcid.org/0000-0003-1197-4081 Silvia Del Vecchio © https://orcid.org/0000-0001-8458-0433 Luana Francesconi © https://orcid.org/0000-0003-1745-7069 Paolo Giordani © https://orcid.org/0000-0003-0087-7315 Pier Luigi Nimis © https://orcid.org/0000-0003-3523-0183 Helmut Mayrhofer © https://orcid.org/0000-0002-9433-3847 Data availability All of the data that support the findings of this study are available in the main text or Supplementary Information. References Altermann S, Leavitt SD, Goward T (2016) Tidying up the genus Letharia: Introducing L. lupina sp. nov. and a new circumscription for L. columbiana. 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Biodiversity and Conservation 30: 2565-2579. https://doi.org/10.1007/s10531-021-02211-w MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 251 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Trontelj P FiSer C (2008) Perspectives: Cryptic species diversity should not be trivialised. Systematics and Biodiversity 7(1): 1-3. https://doi.org/10.1017/ $1477200008002909 White TJ, Bruns T, Lee S, Taylor JW (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ (Eds) PCR Protocols: a Guide to Methods and Applications. Academic Press, New York, 315-322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1 Wirtz N, Printzen C, Lumbsch HT (2008) The delimitation of Antarctic and bipolar spe- cies of neuropogonoid Usnea (Ascomycota, Lecanorales): A cohesion approach of species recognition for the Usnea perpusilla complex. Mycological Research 112: 472-484. https://doi.org/10.1016/j.mycres.2007.05.006 Zanfrognini C (1902) Contribuzione alla flora lichenologica dell’Emilia. Nuovo Giornale Botanico Italiano 9(2): 190-210; (3): 355-377; (4): 434-459. Supplementary material 1 Specimens list, occurence and data Authors: Gabriele Gheza, Chiara Vallese, Luca Di Nuzzo, Simona Corneti, Renato Bene- speri, Elisabetta Bianchi, Giulia Canali, Silvia Del Vecchio, Luana Francesconi, Paolo Giordani, Pier Luigi Nimis, Walter Obermayer, Chiara Pistocchi, Helmut Mayrhofer, Juri Nascimbene Data type: xlsx Explanation note: For each sample, the place and data of collection are provided with coordinates and altitude, the name of the person who collected and identified it, the place where the sample is located (herbarium), chemistry. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.120.154233.suppl1 Supplementary material 2 List of collection sites Authors: Gabriele Gheza, Chiara Vallese, Luca Di Nuzzo, Simona Corneti, Renato Bene- speri, Elisabetta Bianchi, Giulia Canali, Silvia Del Vecchio, Luana Francesconi, Paolo Giordani, Pier Luigi Nimis, Walter Obermayer, Chiara Pistocchi, Helmut Mayrhofer, Juri Nascimbene Data type: xlsx Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.120.154233.suppl2 MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 252 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Supplementary material 3 List of the subset of samples subjected to phylogenetic analysis Authors: Gabriele Gheza, Chiara Vallese, Luca Di Nuzzo, Simona Corneti, Renato Bene- speri, Elisabetta Bianchi, Giulia Canali, Silvia Del Vecchio, Luana Francesconi, Paolo Giordani, Pier Luigi Nimis, Walter Obermayer, Chiara Pistocchi, Helmut Mayrhofer, Juri Nascimbene Data type: docx Explanation note: Data on each sample's collection date, place, and storage is reported, along with the lab's assigned code and markers used. Species names, determined through morphological and chemical analyses and BLAST results, are also provid- ed, with BLAST-based identifications highlighted in bold. Samples marked with an asterisk (*) indicate cases where the phylogenetic analysis results differ from the morphological and chemical findings.Please note that (+) will soon be replaced with GenBank accession numbers. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.120.154233.suppl3 Supplementary material 4 Phylogenetic tree Authors: Gabriele Gheza, Chiara Vallese, Luca Di Nuzzo, Simona Corneti, Renato Bene- speri, Elisabetta Bianchi, Giulia Canali, Silvia Del Vecchio, Luana Francesconi, Paolo Giordani, Pier Luigi Nimis, Walter Obermayer, Chiara Pistocchi, Helmut Mayrhofer, Juri Nascimbene Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.120.154233.suppl4 MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 253 Gabriele Gheza et al.: Revision of the genus Cetrelia in Italy Supplementary material 5 Macrolichens of conservation interest co-occurring with Cetrelia Authors: Gabriele Gheza, Chiara Vallese, Luca Di Nuzzo, Simona Corneti, Renato Bene- speri, Elisabetta Bianchi, Giulia Canali, Silvia Del Vecchio, Luana Francesconi, Paolo Giordani, Pier Luigi Nimis, Walter Obermayer, Chiara Pistocchi, Helmut Mayrhofer, Juri Nascimbene Data type: doc Explanation note: Macrolichens of conservation interest co-occurring with the four Ce- trelia species in the only three Italian sites hosting all of them. Nomenclature follows Nimis and Martellos (2025). Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/mycokeys.120.154233.suppl5 MycoKeys 120: 231-254 (2025), DOI: 10.3897/mycokeys.120.154233 954